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The Cerebellum
• The cerebellum is located dorsal to the pons and medulla under the occipital lobe of the cerebral hemispheres
Cerebellum
The Cerebellum• It is separated from
the occipital lobe by the transverse fissure
• It rests in the posterior cranial fossa of the skull
Cerebellum: Gross Anatomy•It consists of cerebellar cortex and and deep cerebellar nuclei, with white medulla in between
•Cerebellum is uninterrupted across the midline
•The cortex consists of ridges called folia.
•The cerebellum consists of three parts:
•A small inferior part …Flocculonodular lobe (floccular, means a tuft of wool);
•A narrow central vermis (worm shaped)
•Two large lateral hemispheres
nodulus
flocculus
tonsil
Ventral View
Anatomy of the CerebellumAnatomy of the Cerebellum2 symmetrical hemispheres connected medially by the Vermis
Folia: Transversely oriented gyri
3 lobes in each hemisphere: Anterior, Posterior, Flocculonodular (FN)
Neural arrangement: Gray matter (Cortex), White matter (Internal), Scattered cerebellar nuclei: dentate, globose, emboliform, fastigial
Arbor vitae (tree of life): distinctive treelike pattern of the white matter
FoliumFolium
Divisions of the Cerebellum
nodulus
flocculus
Ant LobeAnt Lobe
Ant LobeAnt Lobe
Post LobePost Lobe Post LobePost Lobetonsil
Ventral View Superior Surface
Ant Lobe
Post Lobe
nodulusflocculus
Schematic
vermis
hemisphere
CerebellumCerebellum
Regulation of muscle tone,coordination of skilled voluntary movement
Planning of voluntary activity
Maintenance of balance, control of eye movements
Vestibulocerebellum
Spinocerebellum
Cerebrocerebelum
Anterior Lobe
Posterior Lobe
Flocculo-Nodular Lobe (FN lobe)
Folia
Primary fissure
Cerebellar Subdivisions
•Multiple nomenclatures have been used to describe the various lobules
•Larsell (1952) used Roman Numerals I-X (anterior to posterior)
Archicerebellum-vestibulocerebellum
Paleocerebellum-spinocerebellum
Neocerebellum-pontocerebellum/cerebrocerebellum
Cerebellar Peduncles
MedullaInferior(ICP)
PonsMiddle(MCP)
MidbrainSuperior(SCP)
Connectsto
Peduncle
SCPSCP
SCPSCP SCPSCP
MCPMCP
MCPMCP
MCPMCP
ICPICP
ICPICP
ICPICP
The Macroscopic Anatomy of the Cerebellum
• Gross anatomical divisions– Anterior lobe– Posterior lobe– Flocculonodular lobe
The Evolutionary & Functional Division of the Cerebellum
• The cerebellum can be divided by – Evolutionary age – Function
Cerebellar Subdivisions
•Multiple nomenclatures have been used to describe the various lobules
•Larsell (1952) used Roman Numerals I-X (anterior to posterior)
Archicerebellum-vestibulocerebellum
Paleocerebellum-spinocerebellum
Neocerebellum-pontocerebellum/cerebrocerebellum
The Evolutionary Division of the Cerebellum
• Three phylogenetic divisions within the cerebellum – according to evolutionary age– The flocculonodular lobes (archicerebellum)– The anterior lobes (paleocerebellum)– The posterior lobes (neocerebellum)
Functional Division of the Cerebellum
• Three functional divisions run perpendicular to the phylogenetical divisions
• Align from the midline outwards toward the sides of the body
The Archicerebellum
• Associated with the flocculonodular lobe
• Functions– Balance (vestibular function)
• Receives input from the inferior and medial vestibular nuclei
• Sends fibers back to the vestibular nuclei • Creates a feedback loop that allows for the
constant maintenance of balance
– Eye movement
The Paleocerebellum
• Separated from the anterior lobes by the primary fissure
• Separated from the flocculonodular lobes by the posterolateral fissure
The Paleocerebellum
• Function– Controls proprioception related to muscle tone
• Receives inputs from muscle stretch receptors via the inferior olive (in the medulla)
– Send out put to the inferior cerebellar peduncle
• Receives inputs from the dorsal and ventral spinocerebellar tracts
– Carries information about the position and forces acting on the legs
– Sends axonal projections to the deep cerebellar nuclei
The Neocerebellum
• Receives input from the pontocerebellar tract– From the cerebral cortex (motor) via the pontine
nuclei in the base of the skull
• Axons project to the deep cerebellar nuclei through the middle cerebellar peduncles
• The major output tract of the cerebellum is the superior cerebellar peduncle – Sends signals to the motor cortex and the
supplementary motor area
The Neocerebellum
• Function– Control motor function
• Coordinate fine finger movements
– Feed-forward– Feed-back
Longitudinal Cerebellar Regions
• Vermis– Contributes to body posture
• Paravermal region– Regulates movements of ipsilateral
extremities (e.g. walking)
• Lateral Zone– Regulates skilled movements of ipsilateral
extremity (e.g. tying your shoe)
Deep Cerebellar nucleiDeep Cerebellar nuclei
• Dentate• Emboliform• Globose• Fastigial
Interposed
Fast G E Dent
Simplified Cerebellar Circuitry
Spinal Cord, Cerebral Cortex (via pontine nuclei), Vestibular
SystemCerebellar Cortex
Inferior OliveDeep Cerebellar Nuclei
Brainstem, thalamus, midbrain
Mossy FibersMossy Fibers
Climbing FibersClimbing Fibers
Spinal cord, Cerebral Cortex
+
-
+
INPUT
Microscopic Sections
Outer Molecular layer Stellate cells, Basket cells
Middle Purkinje cell layer Purkinje cells
Inner Granular layer Granule cells, Golgi cells
Cortex
Mol Gr
WM
PurkinjePurkinje
MolecularMolecular
GranularGranular
Major Cell Types
•Granule cells
•Purkinje cells
•Golgi cells
•Stellate cells
•Basket cells
Glutamate
GABA
GABA
GABA
GABA
***The Purkinje cell is the only output of the cerebellar cortex
5 cell types
Stellate (inhibitory)Basket (inhibitory)(molecular layer)
Purkinje (inhibitory)(Purkinje layer)
Golgi (inhibitory)Granule (excitatory)(Granular layer)
Microcircuitry of cerebellum
Inputs:
• Climbing fiber (“+”, excitatory, from inferior olive nucleus)
• Mossy fiber (+, from spinal cord & brain stem)
Output:
• Purkinje cell axon (“-”, inhibitory)
Internal circuitry A. Organization of
cortex is uniform across different subdivisions
B. AFFERENT pathways to the cerebellar cortex excite Purkinje cells.
C. Basket, stellate and Golgi cells regulate Purkinje cell activity.
D. EFFERENT pathways from the cortex originate from Purkinje cells.
Purkinje Cell
Recieves + inputs from parallel fibers and climbing fibers
Recieves - inputs from basket cells, Golgi cells, stellate cells
Heavily invested with glial processes
First direct pathway:
• climbing fiber (+) Purkinje cells (-) deep nuclei
• each climbing fiber projects to 1-10 Purkinje cells
• each Purkinje cell receives input from a single climbing fiber
• Powerful excitatory connection, each climbing fiber spike cause a burst of spikes in Purkinje cell (called a “complex spike”)
complex spike
Second direct pathway:
• Mossy fiber (+) granule cells (axon: parallel fibers, +) Purkinje cells (-) deep nuclei
• each parallel fiber projects to thousands of Purkinje cells (high divergence)
• each Purkinje cell receives input from ~200,000 parallel fibers (high convergence)
• Weak excitatory connection, spatiotemporal summation of inputs from many parallel fibers causes a single spike in Purkinje cell (called a “simple spike”)
glomeruli
simple spikes
Inhibition: focusing in time and space
•Stellate, Golgi, basket cells also receive parallel fiber input and elaborate their dendrites in the molecular layer
•Golgi cells project to granule cells
•Feed forward inhibition
•Stellate, basket cells tend to project laterally
•Collaterals of Purkinje cells also contact nearby cells
+_ _
Lateral inhibition:
• granule cells (axon: parallel fibers, +) stellate and basket cells (-) Purkinje cells in a different row
Negative feedback:
• granule cells (axon: parallel fibers, +) Golgi cells (-) granule cells
Cerebellar PedunclesCerebellar Peduncles
Superior peduncles (to the midbrain):
Fibers originate from neurons in the deep cerebellar nuclei & communicates with the motor cortex via the midbrain and the diencephalon (thalamus)
Middle peduncles (to the pons):
Cerebellum receives information advising it of voluntary motor activities initiated by motor cortex
Inferior peduncles (to the medulla):
Afferents conveying sensory information from muscle proprioceptors throughout the body & from the vestibular nuclei of the brainstem (Spinal cord)
Somatotopic Organization• Tactile information
– Ipsilateral anterior lobule – Bilateral paramedian
lobules– Cerebral Cortex and
Cerebellum have similar representations
• Motor representation– Same area as sensory
mapping– May have auditory and
visual processing
The Cerebellum
• Virtually all fibers entering and leaving the cerebellum are ipsilateral; from and to the same side of the body
Afferent ConnectionsAfferent Connections (1): (1):
1. Inferior Cerebellar Peduncle1. Inferior Cerebellar Peduncle
Restiform BodyRestiform Body Posterior Spinocerebellar TractPosterior Spinocerebellar Tract Olivocerebellar tractOlivocerebellar tract Cuneocerebellar TractCuneocerebellar Tract Reticulocerebellar TractReticulocerebellar Tract
Juxtarestiform BodyJuxtarestiform Body Vestibulocerebellar TractVestibulocerebellar Tract
Cerebellum Cerebellum ConnectionsConnections
Cerebellum Cerebellum ConnectionsConnections
Afferent ConnectionsAfferent Connections (2): (2):
2. Middle Cerebellar Peduncle2. Middle Cerebellar Peduncle
Pontocerebellar fiberPontocerebellar fiber Fibers from raphe nuclei Fibers from raphe nuclei
3. Superior Cerebellar Peduncle3. Superior Cerebellar Peduncle
Anterior Spinocerebellar TractAnterior Spinocerebellar Tract Tecto-cerebellar tract Tecto-cerebellar tract Trigemino-cerebellar tract Trigemino-cerebellar tract
Fibres from locus coeruleusFibres from locus coeruleus
Cerebellum Cerebellum ConnectionsConnections
Cerebellum Cerebellum ConnectionsConnections
Efferent ConnectionsEfferent Connections : :
1. Superior Cerebellar Peduncle1. Superior Cerebellar Peduncle
Cerebellothalamic fiberCerebellothalamic fiber - from 3 deep nuclei to VPLo, VLc, CL- from 3 deep nuclei to VPLo, VLc, CL Cerebellorubral fiberCerebellorubral fiber - from nucleus interpositus - from nucleus interpositus and dentate nucleusand dentate nucleus both goes to cortexboth goes to cortex Cerebello-rubro-spinal tract Cerebello-rubro-spinal tract Fibers to reticular formation and olivary complex Fibers to reticular formation and olivary complex 2. Inferior Cerebellar Peduncle2. Inferior Cerebellar Peduncle
Fastigiovestibular fiberFastigiovestibular fiber Cerebello-reticular Cerebello-reticular cerebello- olivary tract cerebello- olivary tract
Cerebellum Cerebellum ConnectionsConnections
Cerebellum Cerebellum ConnectionsConnections
Outputs of the CerebellumOutputs of the Cerebellum
Dentate nuclei: project contralaterally through the superior cerebellar peduncle to neurons in the contralateral thalamus & from thalamus to motor cortexFunc.: influence planning and initiation of voluntary movement
Emboliform & Globose nuclei: project mainly to the contralateral red nuclei & a small group is projected to the motor cortex Red Nuclei Rubrospinal Tract control of proximal limb muscles
Fastigial nuclei: project to the vestibular nuclei & to the pontine and medullary reticular formation Vestibulospinal & Reticulospinal tracts
Cerebellar nuclei: dentate, globose, emboliform, fastigial
MLFMLF
Main Connections of the VestibulocerebellumMain Connections of the VestibulocerebellumMain Connections of the VestibulocerebellumMain Connections of the Vestibulocerebellum
lower motor neuronlower motor neuron
LMNLMN
vestibulospinal tractvestibulospinal tract
FASTIGIALFASTIGIAL NUCLEUSNUCLEUS
VestibularVestibular OrganOrgan FloculonodularFloculonodular
LobeLobe
VermisVermis
ARCHICEREBELLUMARCHICEREBELLUM
VESTIBULAR NUCLEUSVESTIBULAR NUCLEUS
Main Connections of the PaleocerebellumMain Connections of the PaleocerebellumMain Connections of the PaleocerebellumMain Connections of the Paleocerebellum
lower motor neuronlower motor neuron
SPINAL CORDSPINAL CORD
rubrospinal rubrospinal tracttract
NUCLEUSNUCLEUSINTERPOSITUSINTERPOSITUS
InferiorInferior OlivryOlivryNucleusNucleus
ANTERIOR ANTERIOR LOBELOBEPARAVERMAL PARAVERMAL ZONEZONE
PALEOCEREBELLUMPALEOCEREBELLUM
RED RED NUCLEUSNUCLEUS
spinocerebellar spinocerebellar tracttract
CEREBRALCEREBRAL CORTEXCORTEX
CEREBRALCEREBRAL CORTEXCORTEX
DENTATEDENTATENUCLEUSNUCLEUS
DENTATEDENTATENUCLEUSNUCLEUS
Main Connections of the NeocerebellumMain Connections of the NeocerebellumMain Connections of the NeocerebellumMain Connections of the Neocerebellum
lower motor neuronlower motor neuron
LMNLMN
pyramidal pyramidal tracttract POSTERIOR POSTERIOR
LOBELOBECEREBELLARCEREBELLAR HEMISPHEREHEMISPHERE
POSTERIOR POSTERIOR LOBELOBE
CEREBELLARCEREBELLAR HEMISPHEREHEMISPHERE
THALAMUSTHALAMUSTHALAMUSTHALAMUS
NEOCEREBELLUMNEOCEREBELLUM
PontinePontineNucleusNucleusPontinePontineNucleusNucleus
upper motor neuronupper motor neuron
UMNUMN
upper motor neuronupper motor neuron
UMNUMN
BASALBASALGANGLIAGANGLIABASALBASAL
GANGLIAGANGLIA
Pyramidal Tract and Associated CircuitsPyramidal Tract and Associated CircuitsPyramidal Tract and Associated CircuitsPyramidal Tract and Associated Circuits
lower motor neuronlower motor neuron
UMNUMN
lower motor neuronlower motor neuron
UMNUMN
pyramidal tractpyramidal tract
CerebellumCerebellumCerebellumCerebellum
ReticularFormation
ReticularFormation
CEREBELLUMCEREBELLUM
Cerebellum and Automatic Motor ControlCerebellum and Automatic Motor ControlCerebellum and Automatic Motor ControlCerebellum and Automatic Motor Control
Lower Motor Neuron (LMN)Lower Motor Neuron (LMN)Lower Motor Neuron (LMN)Lower Motor Neuron (LMN)
Motor CortexMotor Cortex
Red NucleusRed Nucleus
Vestibular Nucleus
Vestibular Nucleus
ProprioceptorsProprioceptorsProprioceptorsProprioceptors
Olivocerebellar ConnectionsOlivocerebellar Connections
Caudal portion ofCaudal portion of medial and dorsal accessory olivary nucleusmedial and dorsal accessory olivary nucleus ----------------- vermis of cerebellar cortex (A and B)----------------- vermis of cerebellar cortex (A and B) fastigial nucleusfastigial nucleus vestibular nucleusvestibular nucleus
Rostral portion ofRostral portion of medial and dorsal accessory olivary nucleusmedial and dorsal accessory olivary nucleus ----------------- paravermal region (C----------------- paravermal region (C11, C, C22, C, C33)) nucleus interpositusnucleus interpositus
Principal Inferior Olivary NucleusPrincipal Inferior Olivary Nucleus
----------------- cerebellar hemisphere (D----------------- cerebellar hemisphere (D11, D, D22)) dentate nucleusdentate nucleus
Cerebellum Cerebellum ConnectionsConnections
Cerebellum Cerebellum ConnectionsConnections
CerebellumCerebellum FunctionFunction
CerebellumCerebellum FunctionFunction
Maintenance of EquilibriumMaintenance of Equilibrium - balance, posture, eye movement - balance, posture, eye movement
Coordination of half-automatic movement ofCoordination of half-automatic movement of walking and posture maintenancewalking and posture maintenance - posture, gait - posture, gait
Adjustment of Muscle ToneAdjustment of Muscle Tone
Motor Learning – Motor SkillsMotor Learning – Motor Skills
Cognitive FunctionCognitive Function
Cerebellum: Control of Voluntary MovementCerebellum: Control of Voluntary MovementAll three lobes of cerebellum work together- Comparator of a servo-mechanism All three lobes of cerebellum work together- Comparator of a servo-mechanism
Primary function: 1. To supplement & correlate the activities of other motor areas2. Control of posture 3. Correction of rapid movements initiated by cerebral cortex4. Motor learning
Frequency of nerve impulses in the climbing fibers almost doubles when a monkey learns a new task
Movement Control:a. Inputs from motor cortex inform the cerebellum of an intended
movement before it is initiatedb. Sensory information is then received via the
spinocerebellar tractc. An error signal is generated and is fed back to the cortex
Cerebellum and Motor LearningCerebellum and Motor Learning• Deficits in learning complex motor tasks after
cerebellar lesions• fMRI studies : cerebellum active during learning
of novel movements• Postulated that cerebellar nuclei store certain
motor memories• May be involved in cognitive functionsCerebellar Cognitive FunctionCerebellar Cognitive Function• Plays a role in language and problem solvingPlays a role in language and problem solving• Recognizes and predicts sequences of eventsRecognizes and predicts sequences of events
Cerebellar Processing - 1• The frontal motor association areas of the
cerebral cortex indicates its intents to initiate voluntary muscle contractions
• Through collateral fibers of the pyramdial tracts, it notifies the cerebellum of its activity
Cerebellar Processing - 2• At the same time, the cerebellum receives
information from the proprioceptors throughout the body – Tension in muscles, tendons, and joint positions– From visual and equilibrium pathways
• This information enables the cerebellum to determine where the body is and where it is going – More specifically where the parts of the body
are located in space and how are they moving
Cerebellar Processing - 3• The cerebellar cortex assesses this
information and calculates the best way to coordinate the force, direction, and extent of muscle contraction– Prevents overshoot– Maintains posture– Ensures smooth, coordinated movements
Cerebellar Processing - 4• Via the superior peduncles, the cerebellum
dispatches its “blueprint” for coordination to the cerebral motor cortex which makes appropriate adjustments in its motor plan
• Cerebellar fibers also flow to brain stem nuclei, such as the red nuclei of the midbrain, which in turn project to motor neurons of the spinal cord
The Cerebellum
• The cerebellum continually compares the higher brain’s intention with the body’s performance and sends out messages to initiate the appropriate measures
• In this way, it helps to promote smooth voluntary movements that are precise and economical in terms of muscular effort
The Cerebellum
• Cerebellar injury results in the loss of muscle tone and clumsy, unsure movements, and sometimes even impaired thoughts about movements
Clinical Considerations• Signs of Dysfunction
– Impaired Muscle Synergy– Reduced Muscle Tone– Evident in Skilled Tasks– Nystagmus – Ataxia : incoordination of movement
- decomposition of movement - dysmetria, past-pointing
- dysdiadochokinesia - gait ataxia, truncal ataxia
• Slow Movement (Bradykinesia)• Mild Muscular Weakness (Asthenia)• Asynergia• Speech difficulties (Ataxic Dysarthria)
– affects respiration, phonation, resonance and articulation, but most pronounced in articulation and prosody.
Clinical Considerations 2
• Dysdiadochokinesia– Clumsiness in Alternating Movements– Tapping, Speech Sound
• Dysarthria– Ataxic Dysarthria– Scanning Speech – Slurred and Disjointed Speech
• Dysmetria– Error in Judgment of Range and Distance of Target– Undershooting or Overshooting
Clinical Considerations 3
• Intentional Tremor– Accessory Movement During Volitional Task– vs. Parkinson’s Disease Where Tremor Lessens
During Volitional Movement
• Hypotonia– Reduced Resistance to Passive Stretch
• Rebounding– Inability to Predict Movement – Cannot Hold Back Movement
• Disequilibrium– Unsteady Gait, Body Wavering
Cerebellar Pathologies
• Cerebrovascular Accident (CVA)– Thrombotic, embolic or hemorrhagic– Vertebrobasilar Artery
• Toxicity– Chronic Alcoholism
• Progressive Cerebellar Degeneration– Friedrich's Ataxia: Autosomal Recessive
Heredity Degenerative Condition– Combined Sensory and Motor Dysfunctions
PosturePostureGait – AtaxiaGait – Ataxia
TremorTremor
Cerebellar Cerebellar AtaxiaAtaxia
Ataxic gait and Ataxic gait and position: position: Left cerebellar tumorLeft cerebellar tumor
a. Sways to the right ina. Sways to the right in standing positionstanding position
b. Steady on the b. Steady on the right legright leg
c. Unsteady on the c. Unsteady on the left legleft leg
d. ataxic gaitd. ataxic gait
a b c
d
Cerebellar tumors on Cerebellar tumors on vermisvermis
- Truncal Ataxia- Truncal Ataxia
- Frequent Falling- Frequent Falling
The child in this picture:The child in this picture:
- would not try to stand - would not try to stand unsupportedunsupported
- would not let go of the bed - would not let go of the bed
railrail if she was stood on the floor.if she was stood on the floor.
Cerebellar Cerebellar MedulloblastomaMedulloblastoma